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Paraquanothrus N. Gen. from Freshwater Rock Pools in the USA, with New Diagnoses of Aquanothrus, Aquanothrinae, and Ameronothridae (Acari, Oribatida)

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Paraquanothrus N. Gen. from Freshwater Rock Pools in the USA, with New Diagnoses of Aquanothrus, Aquanothrinae, and Ameronothridae (Acari, Oribatida) Acarologia A quarterly journal of acarology, since 1959 Publishing on all aspects of the Acari All information: http://www1.montpellier.inra.fr/CBGP/acarologia/ [email protected] Acarologia is proudly non-profit, with no page charges and free open access Please help us maintain this system by encouraging your institutes to subscribe to the print version of the journal and by sending us your high quality research on the Acari. Subscriptions: Year 2020 (Volume 60): 450 € http://www1.montpellier.inra.fr/CBGP/acarologia/subscribe.php Previous volumes (2010-2018): 250 € / year (4 issues) Acarologia, CBGP, CS 30016, 34988 MONTFERRIER-sur-LEZ Cedex, France ISSN 0044-586X (print), ISSN 2107-7207 (electronic) The digitalization of Acarologia papers prior to 2000 was supported by Agropolis Fondation under the reference ID 1500-024 through the « Investissements d’avenir » programme (Labex Agro: ANR-10-LABX-0001-01) Acarologia is under free license and distributed under the terms of the Creative Commons-BY-NC-ND which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author and source are credited. Paraquanothrus n. gen. from freshwater rock pools in the USA, with new diagnoses of Aquanothrus, Aquanothrinae, and Ameronothridae (Acari, Oribatida) Roy A. Nortona , Elizabeth Franklinb a State University of New York, College of Environmental Science and Forestry, Syracuse, New York, 13210, U.S.A. b National Institute for Amazonian Research, Biodiversity Coordination, CEP 69067-375 Manaus, Amazonas, Brazil. ABSTRACT Many taxa of mites inhabit long-lived freshwater environments, but the few known to live in small, ephemeral rock pools (lithotelmata) are brachypyline Oribatida. One of these is in the South African genus Aquanothrus (Ameronothridae). We describe adults and juveniles of two new rock-pool species from the USA and propose the sister-genus Paraquanothrus n. gen. to include them. The type-species, Paraquanothrus grahami n. sp., inhabits shallow weathering-depressions (‘pans’) on barren sandstone in the Colorado Plateau, especially southeastern Utah, where it seems to be an opportunistic grazer on microflora and rotifers. Paraquanothrus spooneri n. sp. inhabits rock pools on granite outcrops, is known only from the type-locality in eastern Georgia and appears to ingest mostly plant fragments. Like Aquanothrus, these mites are active only when free water exists. Paraquanothrus shares multiple apomorphic traits with Aquanothrus, for which a new diagnosis is based on corrected information on the type-species, A. montanus, and two undescribed species (one of which is represented in the paratype series). After reviewing historical concepts of Ameronothridae, we propose a new diagnosis (excluding Podacaridae) and propose a new rank and diagnosis for the subfamily Aquanothrinae, which includes Aquanothrus and Paraquanothrus. Molecular studies that have revealed links among Ameronothroidea, Cymbaeremaeoidea and Licneremaeoidea—in ways that question the monophyly of all three superfamilies—are reviewed, and a preliminary evaluation shows morphology to have a modest level of congruence with these results. Keywords oribatid mite, Colorado Plateau, granite outcrop fauna, lithotelma, Ameronothrus, Chudalupia, Scapheremaeus, Scutoverticidae Received 12 December 2017 Zoobank http://zoobank.org/A9FDC4A9-B70D-4965-9F7E-94813BB2929D Accepted 21 February 2018 Published 01 June 2018 Corresponding author Introduction Roy A. Norton: [email protected] Oribatid mites form a dominant component of the soil fauna, being particularly abundant Academic editor and diverse in complex organic horizons of moist forest floors, where they feed primarily on Ekaterina Sidorchuk decaying plant remains and fungi. But several important higher taxa are rarely represented in such habitats; among them are four nominal superfamilies in the hyporder Brachypylina— DOI Licneremaeoidea, Cymbaeremaeoidea, Ameronothroidea and Limnozetoidea—that have 10.24349/acarologia/20184258 adapted to rather different conditions (Weigmann 2006; Norton and Behan-Pelletier 2009). Copyright Xeric soils, and microhabitats that undergo extreme hydric variation regardless of general Norton R.A. and Franklin E. climate (e.g. arboreal and rock surfaces, exposed mosses and lichens), are typical homes Distributed under Creative Commons CC-BY 4.0 How to cite this article Norton R.A. and Franklin E. (2018), Paraquanothrus n. gen. from freshwater rock pools in the USA, with new diagnoses of Aquanothrus, Aquanothrinae, and Ameronothridae (Acari, Oribatida). Acarologia 58(3): 557-627; DOI 10.24349/acarologia/20184258 for members of the first two superfamilies, which overall could be considered ‘dry-adapted’ or ‘dry-tolerant’ (Norton and Behan-Pelletier 2009). By contrast, many members of the third and fourth superfamilies are ostensibly ‘wet-adapted’ or ‘wet-tolerant’. Limnozetoid species are aquatic or semiaquatic in freshwater habitats, and many ameronothroid species are marine-intertidal (Schulte et al. 1975; Behan-Pelletier and Eamer 2007; Pfingstl 2017). But differences in life-style and niche-adaptation among members of these groups are not always as great as such generalities suggest. Oribatid mites in exposed terrestrial microhabitats—e.g. Scutovertex and Scapheremaeus species living in arboreal or epilithic mosses and lichens—may be inactive during dry periods (Travé 1963a; Walter 1999; Smrž 2002; Colloff 2010). Conceptually, this differs little from ‘aquatic’ species in the ameronothroid genera Aquanothrus and Chudalupia, which become immobile when their temporary rock pool habitat no longer contains water (Engelbrecht 1975; Wallwork 1981). At least one species of Scapheremaeus, S. baylyi Colloff, 2010, appears to have a similar lifestyle in rock pools (Colloff 2010). Even the habitats themselves are not necessarily discrete: for example, another species of Scapheremaeus seems to live only in frequently-dried mosses that grow at the edge of exposed, ephemeral rock pools (Norton et al. 2010). Our main purpose is to describe and discuss two new species inhabiting such rock pools (= lithotelmata, gnammas, pans). These are small, temporary lentic environments formed by the accumulation of rainwater in shallow depressions—typically solution or weathering pits—on rock surfaces. They provide habitat for aquatic arthropods and other small invertebrates capable of surviving extended periods of desiccating conditions (Bayly 1997, Jocqué et al. 2010 and cited references). For reasons discussed below, we propose Paraquanothrus n. gen. to include these new species and consider it the sister-genus of Aquanothrus, the type species of which—A. montanus Engelbrecht, 1975—inhabits rock pools in South Africa (Engelbrecht 1975; Jocqué et al. 2006; Vanschoenwinkel et al. 2009). The two new species are endemic to widely separated locations in the USA that present very different environmental conditions. Paraquanothrus grahami n. sp. is restricted to the arid Colorado Plateau, where it inhabits shallow depressions on exposed sandstone. Paraquanothrus spooneri n. sp. is known only from depressions formed in exposed granite outcrops in the warm-temperate state of Georgia. As with A. montanus, adults and juveniles of both species are active only when immersed in water; they burrow into sediment as it dries and remain immobile until water returns. Mites of both genera have been called ‘cryptobiotic’ in the literature (Vanshoenwinkel et al. 2009; Colloff 2010) but they cannot endure desiccation: they survive by preventing it. Secondary objectives are to: (1) improve knowledge of Aquanothrus montanus by adding ontogenetic data, correcting some errors in the original description, critiquing the supposed high variability of the species, and proposing a new generic diagnosis; (2) discuss the systematic position of Paraquanothrus and Aquanothrus by reviewing and critiquing past concepts of Ameronothridae and recasting Aquanothridae as a subfamily of Ameronothridae; and (3) evaluate morphological support for molecular studies that have brought the monophyly of Ameronothroidea, Cymbaeremaeoidea and Licneremaeoidea into question. Materials and methods Acquisition and deposition of specimens — Active adults and juveniles of the two new species were collected directly from rain-filled rock pools by eyedropper or similar device. Inactive specimens were obtained from dry rock-pools by gently collecting sediment. In most cases, this material was rehydrated with rainwater or spring water in the laboratory, then the revived mites were observed alive under a stereomicroscope and manipulated with a fine brush or collected by pipette. Collection data are given below. Numerous preserved adults and juveniles identified as Aquanothrus montanus were studied, including paratype specimens from each of the three South African sites indicated in the original description (Engelbrecht 1975), as Norton R.A. and Franklin E. (2018), Acarologia 58(3): 557-627; DOI 10.24349/acarologia/20184258 558 well as adults and juveniles from Botswana (Thamaga), a population studied by Jocqué et al. (2006). Preserved adults and nymphs of Chudalupia meridionalis Wallwork, 1981 from the type locality in Western Australia were studied. Specimens of other species discussed below are from the general collection of the first author. Type repositories include the mite collections of: the United States National Museum (USNM), the Smithsonian Institution (Washington, D.C.), currently with the US Department
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